DOCSLIB.ORG

The Membrane Transport System of the Guard Cell and Its Integration for Stomatal Dynamics1[CC-BY]

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
The Membrane Transport System of the Guard Cell and Its Integration for Stomatal Dynamics1[CC-BY] Topical Review The Membrane Transport System of the Guard Cell and Its Integration for Stomatal Dynamics1[CC-BY] Mareike Jezek and Michael R. Blatt* Laboratory of Plant Physiology and Biophysics, University of Glasgow, Glasgow G12 8QQ, United Kingdom ORCID IDs: 0000-0002-7460-0792 (M.J.); 0000-0003-1361-4645 (M.R.B.). Stomatal guard cells are widely recognized as the premier plant cell model for membrane transport, signaling, and homeostasis. This recognition is rooted in half a century of research into ion transport across the plasma and vacuolar membranes of guard cells that drive stomatal movements and the signaling mechanisms that regulate them. Stomatal guard cells surround pores in the epidermis of plant leaves, controlling the aperture of the pore to balance CO2 entry into the leaf for photosynthesis with water loss via transpiration. The position of guard cells in the epidermis is ideally suited for cellular and subcellular research, and their sensitivity to endogenous signals and environmental stimuli makes them a primary target for physiological studies. Stomata underpin the challenges of water availability and crop production that are expected to unfold over the next 20 to 30 years. A quantitative understanding of how ion transport is integrated and controlled is key to meeting these challenges and to engineering guard cells for improved water use efficiency and agricultural yields. Stomata are pores that form across the epidermal cell layer of plant leaves and stems. They connect the inner air space of these organs with the atmosphere, thereby serving as the major route for gaseous exchange, bypassing the otherwise impermeable cuticle that forms on the outer epidermal surface. Stomata respond to environmental and endogenous (chemical and hy- draulic) signals, opening and closing the pore in order to satisfy the needs of the mesophyll cells for CO2 in photosynthesis while limiting water loss via transpira- tion to the atmosphere. In the light, stomata may reduce photosynthetic rates by 50% and more when water supply is limiting (Lawson and Blatt, 2014; Vialet- Chabrand et al., 2017). They have a major impact on global water and carbon cycles. Transpiration by crops has been a key factor in global atmospheric modeling and weather prediction for over a quarter of a century (Beljaars et al., 1996; Berry et al., 2010). Today, stomatal transpiration is widely recognized to lie at the center of the crisis in water availability and crop production now expected over the next 20 to 30 years. Water use around the world has increased 6-fold in the past 100 years, 1 This work was supported by the Biotechnology and Biological Sciences Research Council (grant nos. BB/N01832X/1, BB/L001276/1, BB/M01133X/1, BB/M001601/1, and BB/L019205/1), the Euro- pean Union (grant no. NEURICE 678168), and the Leverhulme Trust, the Royal Society, and the John Simon Guggenheim Memorial Foun- dation to M.R.B. * Address correspondence to [email protected]. M.J. assembled the tables; M.J. and M.R.B. prepared the figures and wrote the article. [CC-BY]Article free via Creative Commons CC-BY 4.0 license. www.plantphysiol.org/cgi/doi/10.1104/pp.16.01949 Ò Plant Physiology , June 2017, Vol. 174, pp. 487–519, www.plantphysiol.org Ó 2017 The author(s). All Rights Reserved. 487 Downloaded from on July 10, 2017 - Published by www.plantphysiol.org Copyright © 2017 American Society of Plant Biologists. All rights reserved. Jezek and Blatt twice as fast as the human population, and is expected transporters, it has been possible to connect gene to to double again before 2030, driven mainly by agricul- function through heterologous expression and analysis ture and irrigation (UNESCO, 2015). Indeed, there are in isolation. This same strategy has been used to dissect some very basic reasons for pursuing an understanding macromolecular protein complexes regulating several of how stomata work. K+ channels (Honsbein et al., 2009; Grefen et al., 2015) Stomata attracted the attention of early microsco- and to reconstruct speculative phosphorylation cas- pists, including Grew (1682), who described stomata as cades (Geiger et al., 2009, 2011). Imaging techniques breathing holes on the surface of plant leaves. de combined with voltage clamp studies have shown how Candolle (1827) first confirmed that stomatal apertures are individual transporters are regulated in vivo by cyto- 2+ 2+ variable, but it was only later that von Mohl (1856) solic free [Ca ] ([Ca ]i) and pH (pHi; Thiel et al., 1993; would appreciate the importance of turgor in driving Grabov and Blatt, 1998; Hamilton et al., 2000; Loro these changes. A number of observations recognizable et al., 2012). Site-directed mutation, complementation today followed the advent of the diffusion porometer studies, and structural analysis have uncovered the that enabled measurement of the resistance of the leaf to molecular mechanics of channel gating (Riedelsberger gaseous flow (Darwin and Pertz, 1911). These included et al., 2010; Lefoulon et al., 2014) and early events of transient movements (Darwin, 1916; Knight, 1916), ABA perception and signaling (Garcia-Mata et al., 2003; midday closure (Loftfield, 1921), and the effects of Melcher et al., 2009; Cutler et al., 2010; Wang et al., 2013; drought (Laidlaw and Knight, 1916). Freudenberger Eisenach and Di Angeli, 2017; Inoue and Kinoshita, (1940) and Heath (1948) showed that CO2 within the 2017). leaf air space was important in regulating aperture, and The actions of other hormones such as auxin (Blatt Wilson (1948) established the importance for stomatal and Thiel, 1994; Lohse and Hedrich, 1995), of light and movements of the vapor pressure difference between CO2 (Negi et al., 2008; Kim et al., 2010; Xue et al., 2011; inside and outside the leaf. Kinoshita, 2017), and of plant pathogens (Melotto et al., Stomata were inextricably bound up with the plant 2008) have not been neglected (Melotto et al., 2017). hormone abscisic acid (ABA) when Wright and Hiron Resolving the interface between transport and carbo- (1969) at Wye College in the United Kingdom and hydrate metabolism remains a major challenge (Wang Mittelheuser and van Steveninck (1969) in the United and Blatt, 2011; Horrer et al., 2016; Griffiths and Males, States discovered ABA to be highly effective in closing 2017; Santelia and Lunn, 2017). Considerable attention, stomata and in the subsequent resistance of the leaf to too, has been drawn in recent years to the unusual wilting. This same period, during the 1960s and 1970s, pattern of stomatal development within the epidermis marked a recognition of ion transport, especially of K+ and to its evolution (McElwain et al., 2005; Bergmann salts, and of solute content contributing to the cell tur- and Sack, 2007; Chen et al., 2017). Thus, present interest gor as a driver behind stomatal movements (Fischer in stomata extends well beyond ion transport and gas and Hsiao, 1968; Humble and Hsiao, 1969). Ironically, exchange. Some of these topics are explored in depth in research on stomatal movements at the time was mo- this Focus Issue, and we direct the reader to the several tivated by interest in the mechanism of opening and by Updates accompanying this article (Brodribb and the new concepts of chemiosmosis (Mitchell, 1969). McAdam, 2017; Chater et al., 2017; Vialet-Chabrand Opening, but not closing, was thought to be active, re- et al., 2017). Nonetheless, in many respects, the focus quiring coordination and energy for transport. Re- has come full circle, returning to issues of membrane search came to focus on stomatal closure only following transport and its control. If we are to use our knowledge MacRobbie’s pioneering radiotracer flux analysis in the of stomata to improve crop resilience and agricultural 1980s. Her studies showed that ion efflux during clo- capacity in marginal areas, then stomatal gas exchange sure is a highly coordinated process (MacRobbie, 1981, (Buckley, 2017; Franks et al., 2017) must be linked to an 1983a). understanding of the mechanics of stomatal ion trans- The last three decades have seen an explosion in re- port and its regulation as a priority for the future. Here, search directed to the mechanics of solute transport and we review the current knowledge of ion transport in its regulation. The majority of this new knowledge stomatal guard cells. We emphasize its dynamics and comes from electrophysiological studies, both voltage coordination, the origins of which often defy intuitive clamp on intact stomatal guard cells and patch clamp understanding yet are critical to any rational efforts on guard cell protoplasts, that allow separate transport toward stomatal engineering, and we stress the im- activities to be identified and characterized. These ef- portance of quantitative functional data that are es- forts have provided an unprecedented depth of quan- sential to realize such efforts. titative information about the kinetics of individual ion transporters, including those of the H+-ATPases, K+, 2 Cl , and Ca2+ channels at the plasma membrane and STOMATAL OPENING several cation- and anion-selective channels at the tonoplast, and about the dynamics of their regulation Stomatal pores form between specialized pairs of (Pandey et al., 2007; Sokolovski and Blatt, 2007; Kim epidermal cells, the guard cells (Fig. 1). Guard cells of et al., 2010; Roelfsema and Hedrich, 2010; Lawson dicotyledonous plants bow apart as they expand, and Blatt, 2014). With the cloning of many of these thereby opening the stomatal pore (Table I). The 488 Plant Physiol. Vol. 174, 2017 Downloaded from on July 10, 2017 - Published by www.plantphysiol.org Copyright © 2017 American Society of Plant Biologists. All rights reserved. Guard Cell Ion Transport malate (Mal), as well as sugars. In closing the pore, guard cells reverse this process by metabolizing these solutes or releasing them to the apoplast. The changes in solute content between the open and closed states 2 are substantial, often exceeding 300 to 400 mosmol L 1 (= 300–400 mosM) on a cell volume basis (Table II), and draw corresponding water fluxes, thereby driving the volume and turgor changes that open and close the pore.
Load more

Recommended publications
  • Summary a Plant Is an Integrated System Which: 1
    Summary A plant is an integrated system which: 1. Obtains water and nutrients from the soil. 2. Transports them 3. Combines the H2O with CO2 to make sugar. 4. Exports sugar to where it’s needed Today, we’ll start to go over how this occurs Transport in Plants – Outline I.I. PlantPlant waterwater needsneeds II.II. TransportTransport ofof waterwater andand mineralsminerals A.A. FromFrom SoilSoil intointo RootsRoots B.B. FromFrom RootsRoots toto leavesleaves C.C. StomataStomata andand transpirationtranspiration WhyWhy dodo plantsplants needneed soso muchmuch water?water? TheThe importanceimportance ofof waterwater potential,potential, pressure,pressure, solutessolutes andand osmosisosmosis inin movingmoving water…water… Transport in Plants 1.1. AnimalsAnimals havehave circulatorycirculatory systems.systems. 2.2. VascularVascular plantsplants havehave oneone wayway systems.systems. Transport in Plants •• OneOne wayway systems:systems: plantsplants needneed aa lotlot moremore waterwater thanthan samesame sizedsized animals.animals. •• AA sunflowersunflower plantplant “drinks”“drinks” andand “perspires”“perspires” 1717 timestimes asas muchmuch asas aa human,human, perper unitunit ofof mass.mass. Transport of water and minerals in Plants WaterWater isis goodgood forfor plants:plants: 1.1. UsedUsed withwith CO2CO2 inin photosynthesisphotosynthesis toto makemake “food”.“food”. 2.2. TheThe “blood”“blood” ofof plantsplants –– circulationcirculation (used(used toto movemove stuffstuff around).around). 3.3. EvaporativeEvaporative coolingcooling.
    [Show full text]
  • Co-Ordination of Signalling Elements in Guard Cell Ion Channel Control
    Journal of Experimental Botany, Vol. 49, Special Issue, pp. 351–360, March 1998 Co-ordination of signalling elements in guard cell ion channel control A. Grabov1 and M.R. Blatt Laboratory of Plant Physiology and Biophysics, Wye College, University of London, Wye, Ashford, Kent TN25 5AH, UK Received 3 November 1997; Accepted 10 November 1997 Abstract ably important for providing a plasticity of cellular response to external and environmental stimuli. Fine regulation of solutes transport across the guard Understanding the interdependence and hierarchy of cell plasma membrane for osmotic modulation is signalling elements now presents a major challenge essential for the maintenance of the proper stomatal for research in plant biology. aperture in response to environmental stimuli. The major osmotica, K+,Cl− and malate are transported Key words: Ion channels, stomatal aperture, signalling through selective ion channels in the plasma mem- pathway, second messengers, guard cells. brane and tonoplast of guard cells. To date, a number of ion channels have been shown to operate in the guard cell plasma membrane: outwardly- and inwardly- Introduction + rectifying K channels (IK,in and IK,out), slowly- and Guard cell control of stomatal aperture is crucial to rapid-activating anion channels, and stretch-activated balancing transpiration stream and gas exchange for non-selective channels. Slow and fast vacuolar chan- photosynthesis. As transpiration and CO2 exchange can nels (SV and FV) and voltage-independent K+-selective not be regulated independently, the demand of photosyn- (VK) channels have been found at the guard cell tono- thetic machinery can lead to excessive water evaporation plast. On the molecular level, the regulation of the under adverse environmental conditions.
    [Show full text]
  • Plant Adaptation to Fluctuating Environment and Biomass Production Are Strongly Dependent on Guard Cell Potassium Channels
    Plant adaptation to fluctuating environment and biomass production are strongly dependent on guard cell potassium channels Anne Lebaudy*, Alain Vavasseur†, Eric Hosy*, Ingo Dreyer*‡, Nathalie Leonhardt†, Jean-Baptiste Thibaud*, Anne-Alie´ nor Ve´ ry*, Thierry Simonneau§, and Herve´ Sentenac*¶ *Biochimie et Physiologie Mole´culaire des Plantes, Unite´Mixte de Recherche 5004, Centre National de la Recherche Scientifique/Institut National de la Recherche Agronomique (U.386)/Montpellier SupAgro/Universite´Montpellier 2, 1 Place Viala, 34060 Montpellier Cedex 1, France; †Laboratoire des Echanges Membranaires et Signalisation, Unite´Mixte de Recherche 6191, Centre National de la Recherche Scientifique/Commissariat a`l’Energie Atomique/Universite´ Aix-Marseille, 13108 St. Paul lez Durance Cedex, France; and §Laboratoire d’Ecophysiologie des Plantes sous Stress Environnementaux, Unite´Mixte de Recherche 759, Institut National de la Recherche Agronomique/Montpellier SupAgro, 1 Place Viala, 34060 Montpellier Cedex 1, France Edited by Maarten J. Chrispeels, University of California at San Diego, La Jolla, CA, and approved February 8, 2008 (received for review October 12, 2007) At least four genes encoding plasma membrane inward K؉ chan- plant physiology by engineering an Arabidopsis mutant totally nels (Kin channels) are expressed in Arabidopsis guard cells. A deprived of this activity. double mutant plant was engineered by disruption of a major Kin channel gene and expression of a dominant negative channel Results construct. Using the patch-clamp technique revealed that this Genetic Engineering of an Arabidopsis Mutant Deprived of GCKin mutant was totally deprived of guard cell Kin channel (GCKin) Activity. In a first step, we screened candidate mutant lines in activity, providing a model to investigate the roles of this activity which expression of inward Kϩ channels is affected, by looking in the plant.
    [Show full text]
  • The Number of K Channels in the Plasma Membrane of Guard Cell Protoplasts Changes in Parallel with the Surface Area
    The number of K؉ channels in the plasma membrane of guard cell protoplasts changes in parallel with the surface area Ulrike Homann* and Gerhard Thiel Institute of Botany, Technical University of Darmstadt, D-64287 Darmstadt, Germany Communicated by Enid MacRobbie, University of Cambridge, Cambridge, United Kingdom, May 30, 2002 (received for review August 5, 2001) The activity of the two dominant K؉ channels in the plasma Materials and Methods membrane of Vicia faba guard cell protoplasts was examined .؉ Guard cell protoplasts were prepared from Vicia faba L. cv during pressure-driven swelling. For this purpose, the K currents Bunyan as described (2). For measurements, protoplasts were and the membrane capacitance (C ) of guard cell protoplasts were ͞ ͞ m bathed in 10 mM KCl 10 mM CaCl2 5 mM Mes/KOH, pH 5.6, recorded in parallel. A rise in Cm, reflecting an increase of the and osmolarity was adjusted to 530 mosmol͞kg with sorbitol. membrane surface area, was coupled to a proportional rise in Patch pipettes were filled with 170 mM Kϩ-gluconate͞10 mM ͞ ͞ ͞ ͞ ؉ ؉ conductance of both the K inward and K outward rectifier. The KCl 2 mM MgCl2 2mMMgATP 2 mM EGTA 10 mM Hepes/ ؉ activation kinetics of the K channels were not affected during this KOH, pH 7.8, and osmolarity was adjusted to 560 mosmol͞kg ؉ process. The quantitative and temporal coupling of Cm and K with sorbitol. conductance can hence be interpreted as the result of the addition Whole-cell patch-clamp experiments were performed by using ,of active inward and outward rectifier K؉ channels to the plasma either an EPC-9 patch-clamp amplifier (HEKA Electronics membrane during an increase in surface area.
    [Show full text]
  • Analysis of Guard Cell Viability and Action in Senescing Leaves
    Plant Physiol. (1985) 79, 7- 10 0032-0889/85/79/0007/04/$0 1.00/0 Analysis of Guard Cell Viability and Action in Senescing Leaves of Nicotiana glauca (Graham), Tree Tobacco' Received for publication January 25, 1985 and in revised form April 22, 1985 RICHARD OZUNA, RAMON YERA, KIM ORTEGA, AND GARY TALLMAN* Natural Science Division, Pepperdine University, Malibu, California 90265 ABSTRACT not exhibit signs of functionality (24). As a result, Zeiger and Downloaded from https://academic.oup.com/plphys/article/79/1/7/6081518 by guest on 02 October 2021 Schwartz (24) have suggested that yellowing leaves retain sto- In an attempt to determine whether low epidermal conductances to matal control throughout the senescence process. water vapor diffusion of senescing leaves were caused by internal changes These studies raise the question of how stomatal movements in guard cells or by factors external to guard cells, stomatal behavior was are controlled in senescing leaves. Stomates of senescing leaves examined in intact senescing and nonsenescing leaves ofNicotiana glauca may fail to open fully because properties of guard cells that (Graham), tree tobacco, grown in the field or in an environmental cham- facilitate turgor production are damaged or modified. Alterna- ber. Conductances of senescing leaves were 5 to 10% of the maximum tively, stomatal movements in senescing leaves might be con- conductances of nonsenescing leaves of the same plant, yet guard cell trolled solely by factors external to guard cells. duplexes isolated from epidermal peels of senescing leaves developed full In this paper, we examine the diurnal patterns oftranspiration turgor in the light in solutions containing KCI, and sodium cobaltinitrite and epidermal conductance, and the water relations of senescing staining showed that K+ accumulated as turgor developed.
    [Show full text]
  • Xylem Sap Ph Increase: a Drought Signal Received at The
    Plant Physiol. (1 997) 113: 559-573 Xylem Sap pH Increase: A Drought Signal Received at the Apoplastic Face of the Guard Cell That lnvolves the Suppression of Saturable Abscisic Acid Uptake by the Epidermal Symplast' Sally Wilkinson and William J. Davies* Division of Biological Sciences, lnstitute of Environmental and Biological Sciences, Lancaster University, Bailrigg, Lancaster, LAI 4YQ, United Kingdom is synthesized in roots that are in contact with drying soil Drought increased the pH of Commelina communis xylem sap (Davies and Zhang, 1991). The concentration of ABA in the from 6.1 to 6.7. Conductances of transpiring leaves were 50% xylem sap of well-watered sunflower plants collected by lower in pH 7.0 than in pH 6.0 buffers, but bulk leaf abscisic acid using techniques that match the transpirational flow rate of (ABA) concentration and shoot water status were unaffected by pH. intact plants is between 1.0 and 15.0 nmol dm-3 (Schurr et Stomatal apertures of isolated abaxial epidermis incubated on sim- al., 1992). In water-stressed sunflower plants the concen- ple buffers increased with external pH, so in vivo this must be tration of ABA can reach 3.0 pmol dmP3. Applications of overridden by alternative pH effects. Reductions in leaf transpira- similar concentrations of synthetic ABA to the transpira- tion rate at pH 7.0 were dependent on the presence of 1OP8 mo1 tion stream of detached leaves with normal water poten- dm-3 ABA in the xylem stream. We inferred that at pH 7.0 leaf apoplastic ABA concentrations increased: pH did not affect distri- tials commonly reduce water loss by approximately 50%.
    [Show full text]
  • Polarimetric Imaging Microscopy for Advanced Inspection of Vegetal
    www.nature.com/scientificreports OPEN Polarimetric imaging microscopy for advanced inspection of vegetal tissues Albert Van Eeckhout1*, Enrique Garcia‑Caurel2, Teresa Garnatje3, Juan Carlos Escalera1, Mercè Durfort4, Josep Vidal1, José J. Gil5, Juan Campos1 & Angel Lizana1 Optical microscopy techniques for plant inspection beneft from the fact that at least one of the multiple properties of light (intensity, phase, wavelength, polarization) may be modifed by vegetal tissues. Paradoxically, polarimetric microscopy although being a mature technique in biophotonics, is not so commonly used in botany. Importantly, only specifc polarimetric observables, as birefringence or dichroism, have some presence in botany studies, and other relevant metrics, as those based on depolarization, are underused. We present a versatile method, based on a representative selection of polarimetric observables, to obtain and to analyse images of plants which bring signifcant information about their structure and/or the spatial organization of their constituents (cells, organelles, among other structures). We provide a thorough analysis of polarimetric microscopy images of sections of plant leaves which are compared with those obtained by other commonly used microscopy techniques in plant biology. Our results show the interest of polarimetric microscopy for plant inspection, as it is non‑destructive technique, highly competitive in economical and time consumption, and providing advantages compared to standard non‑polarizing techniques. Te inherent properties of light are a signifcant source of information when used to probe the properties of vegetal tissues1–17. In microscopy imaging, it is common to prepare the samples to be studied in very thin sec- tions to prevent multiple scattering of light by the diferent tissue structures, which can degrade image contrast and spatial resolution.
    [Show full text]
  • Water Movement in Plants
    Chapter 3 - Water movement in plants Chapter editors: Brendan Choat and Rana Munns Contributing Authors: B Choat1, R Munns2,3,4, M McCully2, JB Passioura2, SD Tyerman4,5, H Bramley6 and M Canny* 1Hawkesbury Institute of the Environment, University of Western Sydney; 2CSIRO Agriculture, Canberra; 3School of Plant Biology, University of Western Australia; 4ARC Centre of Excellence in Plant Energy Biology; 5School of Agriculture, Food and Wine, University of Adelaide; 6Facutly of Agriculture and Environment, University of Sydney; *Martin Canny passed away in 2013 Contents 3.1 - Plant water relations 3.2 - Long distance xylem transport 3.3 - Leaf vein architecture and anatomy 3.4 - Water movement from soil to roots 3.5 - Water and nutrient transport through roots 3.6 - Membrane transport of water and ions 3.7 - References Evolutionary changes were necessary for plants to inhabit land. Aquatic plants obtain all their resources from the surrounding water, whereas terrestrial plants are nourished from the soil and the atmosphere. Roots growing into soil absorb water and nutrients, while leaves, supported by a stem superstructure in the aerial environment, intercept sunlight and CO2 for photosynthesis. This division of labour results in assimilatory organs of land plants being nutritionally inter-dependent; roots depend on a supply of photoassimilates from leaves, while shoots (leaves, stems, flowers and fruits) depend on roots to supply water and mineral nutrients. Long-distance transport is therefore a special property of land plants. In extreme cases, sap must move up to 100 m vertically and overcome gravity to rise to tree tops. Karri (Eucalyptus diversicolor) forest in Pemberton, Western Australia.
    [Show full text]
  • Exchange & Transport in Plants Heyer 1
    Exchange & Transport in Plants Exchange & Transport Plant Exchange & Transport in Vascular Plants I. Water and Solute Uptake by Cells II. Local Transport III. Long Distance Transport IV. Gas Exchange Passive Transport (Diffusion) Water and Solutes - Uptake by Cells • Net movement of molecules from a region of high concentration to a region of low concentration Caused by random (Brownian) movements of molecules (Increase entropy) Each type of molecule follows its own concentration gradient At equilibrium, movement is equal in both directions Osmosis: simple diffusion of the solvent (water) Water and Solute Uptake by Cells Water moves across lipid bilayer and through Osmotic pressure aquaporins (membrane gate proteins) • Water diffuses Move the water by moving the solutes! according to its . Hypertonic solution: higher concentration of concentration gradient solutes • ↑Osm → ↓[water] ↓ Osm →↑[water] . Hypotonic solution: lower concentration of • Osmosis can solutes generate force (osmotic pressure) . Isotonic solution: equal solute concentrations Semipermeable membrane Heyer 1 Exchange & Transport in Plants Water and Solute Uptake by Cells Water and Solute Uptake by Cells Hypotonic Isotonic Hypertonic solution solution solution Plant cells flaccid Animal cell Plant cell Plant cells turgid Turgor pressure: pressure exerted on wall of turgid cell = 75–100psi Water Potential (Ψ) Water Potential (Ψ) • Osmotic pressure pulls water to the • Physical pressure • Pure water, ΨS = 0 MPa. • pressure potential [ΨP] could right. (= pressure potential [ΨP]): . 1 MPa = 10 atm = ~160 psi be negative solution on right has potential • Osmotic potential • solutes lower energy to push water to the left ΨS (=solute potential [ΨS]): solution on left has potential . 0.1 Osm = –0.23 MPa • If ΨS & ΨP are equal but energy to push water to the right opposite→ no net flow • Water moves from • ↓Osm→↑ ΨS high Ψ to low Ψ.
    [Show full text]
  • Plant Structure and Function 617 DO NOT EDIT--Changes Must Be Made Through “File Info” Correctionkey=A
    DO NOT EDIT--Changes must be made through “File info” CorrectionKey=A Plant Structure CHAPTER 21 and Function Big idea Plants are organisms with specialized cells that absorb light and carry out photosynthesis and tissue systems that absorb, transport, and store water and nutrients. 21.1 Plant Cells and Tissues 5B, 10B, 10C 21.2 The Vascular System 4B, 5B, 10B, 10C 21.3 Roots and Stems 4B, 5B, 10B, 10C data analysis ideNTifyiNg The importaNCe Of RePeaTed TRiaLS 2G 21.4 Leaves 4B, 5B, 10B, 10C Online BiOlOgy HMDScience.com ONLiNe Labs ■■ Virtual Lab Plant Transpiration ■■ Density of Stomata ■■ Open inquiry Lab The Vascular System ■■ QuickLab Chlorophyll Fluorescence ■■ Open inquiry Lab Roots and Stems ■■ Photosynthesis and Red Leaves ■■ Connecting Form to Function ■■ Comparing Plant Structures ■■ Absorption Spectra of Plant Pigments (t) ©José Fuste Raga/Age Fotostock 616 Unit 7: Plants DO NOT EDIT--Changes must be made through “File info” CorrectionKey=A Q how would this tree compete with other species? Fig trees (Ficus) have a unique way of growing. Many trees of this genus are called strangler figs because their aggressive growth actually strangles other trees. Strangler figs can also wrap around unmoving objects such as these temple walls. Their seeds germinate easily in tree branches or building cracks, and then snakelike roots grow down to the ground. R e a d i N g T o o L b o x This reading tool can help you learn the material in the following pages. USiNg LaNgUAGE Your TURN Cause and effect In biological processes, one step leads Identify the cause and the effect in the following sentences.
    [Show full text]
  • Chapter 36 Transport in Vascular Plants Lecture Outline
    Chapter 36 Transport in Vascular Plants Lecture Outline Overview: Pathways for Survival The algal ancestors of plants obtained water, minerals and CO2 from the water in which they were completely immersed. For vascular plants, the evolutionary journey onto land involved the differentiation of the plant body into roots, which absorb water and minerals from the soil, and shoots, which absorb light and atmospheric CO2 for photosynthesis. This morphological solution created a new problem: the need to transport materials between roots and shoots. Xylem transports water and minerals from the roots to the shoots. Phloem transports sugars from the site of production to the regions that need them for growth and metabolism. Concept 36.1 Physical forces drive the transport of materials in plants over a range of distances Transport in plants occurs on three levels: 1. The uptake and loss of water and solutes by individual cells, such as root hairs. 2. Short-distance transport of substances from cell to cell at the level of tissues or organs, such as the loading of sugar from photosynthetic leaf cells into the sieve tubes of phloem. 3. Long-distance transport of sap within xylem and phloem at the level of the whole plant. Transport at the cellular level depends on the selective permeability of membranes. The selective permeability of a plant cell’s plasma membrane controls the movement of solutes between the cell and the extracellular solution. Molecules tend to move down their concentration gradient. Diffusion across a membrane is called passive transport and occurs without the direct expenditure of metabolic energy by the cell.
    [Show full text]
  • Anatomy of Flowering Plants
    84 BIOLOGY CHAPTER 6 ANATOMY OF FLOWERING PLANTS 6.1 The Tissues You can very easily see the structural similarities and variations in the external morphology of the larger living organism, both plants and 6.2 The Tissue animals. Similarly, if we were to study the internal structure, one also System finds several similarities as well as differences. This chapter introduces 6.3 Anatomy of you to the internal structure and functional organisation of higher plants. Dicotyledonous Study of internal structure of plants is called anatomy. Plants have cells and as the basic unit, cells are organised into tissues and in turn the tissues Monocotyledonous are organised into organs. Different organs in a plant show differences in Plants their internal structure. Within angiosperms, the monocots and dicots are also seen to be anatomically different. Internal structures also show 6.4 Secondary adaptations to diverse environments. Growth 6.1 THE TISSUES A tissue is a group of cells having a common origin and usually performing a common function. A plant is made up of different kinds of tissues. Tissues are classified into two main groups, namely, meristematic and permanent tissues based on whether the cells being formed are capable of dividing or not. 6.1.1 Meristematic Tissues Growth in plants is largely restricted to specialised regions of active cell division called meristems (Gk. meristos: divided). Plants have different kinds of meristems. The meristems which occur at the tips of roots and shoots and produce primary tissues are called apical meristems (Figure 6.1). 2021-22 ANATOMY OF FLOWERING PLANTS 85 Central cylinder Cortex Leaf primordium Protoderm Shoot apical Meristematic zone Initials of central cylinder Root apical and cortex Axillary bud meristem Differentiating Initials of vascular tissue root cap Root cap Figure 6.1 Apical meristem: (a) Root (b) Shoot Root apical meristem occupies the tip of a root while the shoot apical meristem occupies the distant most region of the stem axis.
    [Show full text]